18/2 • 2019, 289–312 DOI: 10.2478/hacq-2018-0013 Kenophytes in different forest types of Ukraine Viktor Onyshchenko1 Key words: forest, vegetation, Abstract alien plants, kenophytes, invasion, The article aims to 1) describe the distribution of non-native species of vascular habitats. plants (only kenophytes, i. e. naturalized species introduced after the 15th cen- tury) in different types of forests and different biogeographical regions of Ukraine; Ključne besede: gozd, vegetacija, 2) compare proportions of kenophytes in forests of different areas; 3) detect statis- tujerodne rastline, kenofiti, invazija, tically significant changes in the occurrence of kenophytes over the last 80 years. habitati. The material consists of 2701 relevés sampled in 1990‒2018. They were taken from Ukrainian phytocoenological publications and databases. In Ukraine, as in other European countries, the highest proportion of kenophytes (percentage of species number per relevé) is in floodplain forests (up to 9.1% in willow and pop- lar forests). The lowest proportion is characteristic for bog woods (0%) and most types of mountain forests. Among biogeographical regions, the highest values were found in the flatland areas of the Steppic region and the forest-steppe part of the Continental region. The most frequent species are Impatiens parviflora (predomi- nantly in broadleaved woods, absent in relevés from the steppic biogeoregion) and Erigeron canadensis (pine woods on sand). Comparison with 1466 older relevés sampled in 1950–1989 allows us to make a conclusion that the proportion of kenophytes increased at least in one habitat type (oak-hornbeam forests). Izvleček V članku smo želeli 1) opisati razširjenost tujerodnih rastlinskih vrst (samo keno- fitov, to je naturaliziranih vrst, ki so se pojavile po 15. stoletju) v različnih gozdnih habitatnih tipih in v različnih biogeografskih regijah v Ukrajini; 2) primerjati deleže kenofitov v gozdovih v različnih območjih; 3) prikazati statistično značilne spremembe v pojavljanju kenofitov v zadnjih 80. letih. Podatke predstavlja 2701 popis, ki so bili narejeni v obdobju med 1990 in 2018. Uporabili smo fitoceno- loško literaturo in podatkovne baze iz Ukrajine. V Ukrajini, kot tudi v ostalih državah v Evropi, je najvišji delež kenofitov (odstotek vrst na popis) v poplavnih gozdovih (do 9,1% v vrbovih in topolovih gozdovih). Najnižji odstotek je značilen za barjanske gozdove (0%) in večino habitatnih tipov gorskih gozdov. Med bioge- ografskimi regijami je najvišji delež v nižinskih predelih stepske regije in gozdne stepe v kontinentalni regiji. Najbolj pogosti vrsti sta Impatiens parviflora(prevla - duje v listopadnih gozdovih, odsotna je v popisih iz stepske biogeografske regije) Received: 13. 4. 2018 in Erigeron canadensis (v borovih gozdovih na pesku). S primerjavo 1466 starejših Revision received: 4. 12. 2018 popisov, narejenih med leti 1950 in 1989 smo prišli do zaključka, da se je delež Accepted: 5. 1. 2019 kenofitov povečal vsaj v enem habitatnem tipu (hrastovo-gabrovi gozdovi). 1 M. G. Kholodny Institute of Botany of the National Academy of Sciences of Ukraine, Ukraine. E-mail: [email protected] 289 Viktor Onyshchenko 18/2 • 2019, 289–312 Kenophytes in different forest types of Ukraine ropean Russia occupy 49% of the study area but they are Introduction represented only by 1% of the total number of relevés. Phytosociological relevés are an important source of in- We made the assessment of occurrences of kenophytes in formation about ecological preferences of species, their forests of Ukraine using a higher number of relevés. A geographical distribution and dynamics. In recent decades more narrow geographical scope allows using the results some studies were published that show geographical and for comparisons between regions of Europe. The analy- ecological patterns of alien species on the basis of phyto- sis for the territory of Ukraine, even without compari- coenological data. The study areas were the Czech Repub- son with other countries, may reveal some geographical lic (Chytrý at al. 2005), Catalonia (Vila et al. 2007), the patterns because Ukraine comprises parts of four biogeo- Basque Country (Campos et al. 2013), Slovakia (Med- graphical regions and measures about 1300 km in length. vecká et al. 2014) and Slovenia (Küzmič & Šilc 2017). At present there is no information in literature about the Chytrý et al. (2008) performed a comparison of plant level of invasion in forests of the Steppic biogeographical invasions for three areas of Europe (the Czech Republic, region of Europe. In other bioregions of Europe, estima- Catalonia, Great Britain) that represent three biogeo- tions were carried out only at higher levels of habitat clas- graphical regions (Continental, Mediterranean, Atlantic). sifications. Estimations in the aforementioned articles were made for Presence of 40 alien invasive species within plant com- classes and alliances of vegetation or for EUNIS level 2 munities of Ukraine at the level of alliances of vegetation habitats (Davies et al 2004, EUNIS habitat …). Accord- without quantitative estimations was described in article ing to these studies the lowest proportions of aliens in by Abduloyeva & Karpenko (2009). all regions are in bogs, mires, alpine and subalpine grass- The aim of the present study is to describe the frequency lands, heaths on poor soils. The highest proportions are of kenophytes in forest types and different biogeographi- found in synanthropic vegetation, marine coastal habitats, cal regions of Ukraine, and to compare the results with standing inland waters (only kenophytes), sedge and reed published data on the subject from other areas. beds, riverine shrubs. Wagner et al. (2017) published the assessment of plant invasions in European forests that is based on the analy- Materials and methods sis of 83,396 relevés from the European Vegetation Ar- The study area includes the entire territory of Ukraine. chive (Chytrý at al. 2016) sampled in 1970–2015. The According to the data on the European Environment assessment was made for habitat types of the revised Agency website (Europe’s biodiversity ... 2018) Ukraine version of EUNIS habitat classification (Schaminée et al. comprises parts of four European biogeographical re- 2014). These habitat types approximately correspond to gions: Alpine, Continental, Pannonian and Steppic. Two level 3 or 4 habitats of the official EUNIS habitat classifi- regions (Continental and Steppic) were subdivided ac- cation. All analyses were performed only for kenophytes cording to the geobotanical subdivision of Ukraine in the (naturalized species introduced voluntarily or involun- National Atlas of Ukraine (2008) (Figure 1). tarily after the 15th century). Archaeophytes (introduced The relevés for the analysis were taken from the data- before the end of the 15th century) were excluded from base EU-UA-006 “Vegetation Database of Ukraine and consideration because the status of archaeophytes is poorly known in some countries. The highest proportion of kenophytes was found in riparian woods. The lowest proportions were in mountain forests and mire wood- lands. The most frequent kenophytes of European woods turned out to be Impatiens parviflora (21.1% of total fre- quency of kenophytes), Padus serotina (8.8%), Robinia pseudoacacia (7.5%). The highest number of occurrences of kenophytes belongs to phanerophytes and therophytes (for native species hemicryptophytes and phanerophytes). Species originating in Europe constitute 47% of the to- tal number of kenophytes in European woods, and their total frequency is 22% of total frequency of kenophytes. The distribution of relevés used in the article by Wag- ner et al. (2017) has a strong geographic bias: Ukraine, Figure 1: Biogeographical regions of Ukraine. Belarus, Moldova, Georgia, Armenia, Azerbaijan and Eu- Slika 1: Biogeografske regije Ukrajine. 290 Viktor Onyshchenko 18/2 • 2019, 289–312 Kenophytes in different forest types of Ukraine Adjacent Parts of Russia”, unpublished database of rele- of their natural range or in clearly unnatural conditions vés from the National Nature Park “Holosiivskyi”(Kyiv and natural woods of non-native species), 2) relevés that city) and numerous Ukrainian publications (Andrienko have no species data obtained in period May 15 – Au- et al. 2004, Bairak 1996, Budzhak & Onyshchenko gust 31, 3) relevés that could not be unequivocally as- 2004, Chorney et al. 2005, Derzhypilsky et al. 2011, signed to either habitat type. The area of the relevés varies Didukh 1996, 2003, Didukh et al. 2003, Fitsailo 2003, from 100 to 2500 m2, predominantly 400–900 m2. This Gomlya 2005, Goncharenko 2001, 2003, Klimuk et al. parameter was not used in relevé filtering. Some analy- 2006, Konishchuk 2003, Kuzemko 2001, Kuzemko & ses showed that the effect of plot size on the proportion Chorna 2002, Lukash & Onyshchenko 2006, Melnyk & of species is negligible (Chytrý et al. 2005, Vilà et al. Korinko 2005, Onyshchenko 1998, 2002, 2007, 2009, 2007). The final number of relevés was 2701. In order to 2013a, 2013b, Onyshchenko & Budzhak 2003, Onysh- compare the current frequency of kenophytes with their chenko &t Lubinska 2006, Onyshchenko & Lukash frequency in previous decades 1466 relevés sampled in 2004, Onyshchenko & Sidenko 2002, Onyshchenko & 1950–1989 were used (Bukhalo 1962, Grygora et al. Yuglichek 2010, Onyshchenko et al. 2007, 2009, 2015, 2005, Vorobyov et al. 1997, Vorobyov et al. 2008). Old 2016, Orlov & Yakushenko 2005, Panchenko 2013, relevés were not sampled in the